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Introductory

SPECIES: Gymnocarpium dryopteris | Oak Fern
ABBREVIATION : GYMDRY SYNONYMS : Dryopteris disjuncta (Ledeb.) Mort. Dryopteris linnaeana Christens. Phegopteris dryopteris (L.) Fee Thelypteris dryopteris (L.) Slosson SCS PLANT CODE : GYDR COMMON NAMES : oak fern TAXONOMY : The commonly accepted scientific name for oak fern is Gymnocarpium dryopteris (L.) Newm. in the family Polypodiaceae. There are two subspecies as follows [18]: Gymnocarpium dryopteris ssp. disjunctum (Rupr.) Sarvela G. dryopteris ssp. dryopteris The synonym Dryopteris disjuncta is used frequently in the literature [8,9,22]. LIFE FORM : Fern or Fern Ally FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : S. A. Snyder, August 1993 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Snyder, S. A. 1993. Gymnocarpium dryopteris. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Gymnocarpium dryopteris | Oak Fern
GENERAL DISTRIBUTION : Oak fern has a circumboreal distribution [16]. In North America it occurs form Alaska south to isolated populations in Oregon and east across all provinces of Canada to the Atlantic Coast. It occurs throughout New England south to Virginia and west to Ohio. Scattered populations are found in the northern Midwest states of Wisconsin, Michigan, and Minnesota, and it reaches as far south as Iowa. Oak fern also occurs in isolated populations of the Intermountain West and in New Mexico and Arizona [28]. Gymnocarpium dryopteris ssp. disjunctum is found along the West Coast and in parts of Idaho and eastern Washington [18]. ECOSYSTEMS : FRES11 Spruce - fir FRES18 Maple - beech - birch FRES19 Aspen - birch FRES20 Douglas-fir FRES22 Western white pine FRES23 Fir - spruce FRES24 Hemlock - Sitka spruce FRES25 Larch FRES26 Lodgepole pine FRES28 Western hardwoods STATES : AK AZ CO CT ID IA ME MD MA MI MN MT NH NJ NM NY OH OR PA RI SD VT VA WA WV WI WY AB BC MB NB NF NT NS ON PE PQ SK YT ADMINISTRATIVE UNITS : ACAD APIS BLCA BLRI CUVA DEWA DENA GLBA GLAC GRTE ISRO LACL MORA NOCA OLYM PIRO ROMO SAJH SHEN SLBE VOYA WRST YUCH BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 5 Columbia Plateau 8 Northern Rocky Mountains 11 Southern Rocky Mountains 12 Colorado Plateau KUCHLER PLANT ASSOCIATIONS : K001 Spruce - cedar - hemlock forest K002 Cedar - hemlock - Douglas-fir forest K003 Silver fir - Douglas-fir forest K004 Fir - hemlock forest K005 Mixed conifer forest K008 Lodgepole pine - subalpine forest K012 Douglas-fir forest K013 Cedar - hemlock - pine forest K014 Grand fir - Douglas-fir forest K015 Western spruce - fir forest K020 Spruce - fir - Douglas-fir forest K021 Southwestern spruce - fir forest K093 Great Lakes spruce - fir forest K096 Northeastern spruce - fir forest K099 Maple - basswood forest K102 Beech - maple forest K103 Mixed mesophytic forest K106 Northern hardwoods K107 Northern hardwoods - fir forest K108 Northern hardwoods - spruce forest SAF COVER TYPES : 5 Balsam fir 12 Black spruce 13 Black spruce - tamarack 16 Aspen 18 Paper birch 23 Eastern hemlock 25 Sugar maple - beech - yellow birch 26 Sugar maple - basswood 27 Sugar maple 35 Paper birch - red spruce - balsam fir 37 Northern white-cedar 38 Tamarack 60 Beech - sugar maple 107 White spruce 201 White spruce 202 White spruce - paper birch 203 Balsam poplar 204 Black spruce 205 Mountain hemlock 206 Engelmann spruce - subalpine fir 212 Western larch 213 Grand fir 215 Western white pine 217 Aspen 218 Lodgepole pine 221 Red alder 223 Sitka spruce 224 Western hemlock 225 Western hemlock - Sitka spruce 226 Coastal true fir - hemlock 227 Western redcedar - western hemlock 228 Western redcedar 229 Pacific Douglas-fir 230 Douglas-fir - western hemlock 251 White spruce - aspen 252 Paper birch 253 Black spruce - white spruce 254 Black spruce - paper birch SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Publications listing oak fern as a dominant species are as follows: Preliminary classification of forest vegetation of the Kenai Peninsula, Alaska [30] Forest habitat types of northern Idaho: a second approximation [7] Classification and management of riparian and wetland sites in northwest Montana [4]

VALUE AND USE

SPECIES: Gymnocarpium dryopteris | Oak Fern
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Grizzly bear in the Selkirk Mountains, Idaho, have been observed eating oak fern fronds [1]. Elk on Vancouver Island eat oak fern, but use is low in spring and summer [15]. PALATABILITY : NO-ENTRY NUTRITIONAL VALUE : NO-ENTRY COVER VALUE : NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES : NO-ENTRY OTHER USES AND VALUES : Because of its ease and success at transplanting, oak fern is a desirable garden plant [16]. MANAGEMENT CONSIDERATIONS : Oak fern can interfere with the growth of Engelmann spruce (Picea engelmanni) seedlings [5]. Glyphosate can injur oak fern when applied from July to September [26]. It controls growth of oak fern following harvesting, allowing growth of desired tree species. Oak fern responses to logging vary. In areas where logging leads to decreases in site moisture, oak fern will decrease [8]. In wet, high-elevation areas logging can increase oak fern abundance [25].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Gymnocarpium dryopteris | Oak Fern
GENERAL BOTANICAL CHARACTERISTICS : The deciduous oak fern is delicate in appearance and grows up to 11.8 inches (30 cm) tall [17]. Its petioles are 4 to 12 inches (10-30 cm) long and parallel to the ground [21]. The blade is divided into three triangular leaflets [17]. Each petiole arises from a single node on the creeping rhizome [21]. Spore covers are absent [16]. RAUNKIAER LIFE FORM : Chamaephyte Geophyte REGENERATION PROCESSES : Oak fern reproduces by spores and sprouts from rhizomes. The spores are adapted for high wind dispersal [18]. There is much outcrossing in this species, and no intragametophytic fertilization [18]. Spores have been found in soil seedbanks where adult plants are absent [23]. Spores have sprouted in a greenhouse from soil samples taken from beneath canopy gaps in northern hardwood forests [24]. SITE CHARACTERISTICS : Oak fern occurs on mesic to wet sites in mixed conifer and northern hardwood stands [20]. Soils are moist to well-drained, with pH ranging from 4.5 to 6.4 [7,10,36]. Soil textures are gravelly or sandy to silty clay loams [4,7]. Oak fern grows at elevations from 21 to 1,700 feet (7-518 m) in the Adirondacks [20]. In Alberta it occurs from 1,960 to 4,300 feet (600-1,300 m), and in Idaho oak fern occurs at elevations between 2,500 and 4,500 feet (760-1,370 m) [7,10]. Oak fern occurs on moderately steep slopes and northeast to north and west aspects [7,10]. Some plant species associated with oak fern include Alaska-cedar (Chamaecyparis nootkatensis), devil's club (Oplopanax horridus), alder (Alnus spp.), mountain maple (Acer spicatum), red-osier dogwood (Cornus sericea), twinberry honeysuckle (Lonicera involucrata), thimbleberry (Rubus parviflorus), prickly rose (Rosa acicularis), highbush cranberry (Viburnum edule), twinflower (Linnaea borealis), heartleaf arnica (Arnica cordifolia), starry Solomon's-seal (Smilacina stellata), and bluejoint reedgrass (Calamagrostis canadensis) [7,10,12,14,17,30]. SUCCESSIONAL STATUS : Facultative Seral Species Oak fern is an indicator of cool, moist sites and mid- to late-seral forests [4,21,22,30]. Oak fern will grow on disturbed sites before canopy cover is established in the subboreal spruce (Picea) zone of British Columbia [14]. It is present in that zone in both mesic seral communities and climax forests. Similarly, in spruce-hemlock (Tsuga) forests of southeast Alaska oak fern will begin establishing in 25- to 35-year-old stands following disturbance by logging or fire [1]. They will then dominate the understory for the following century. Oak fern has been used as a site-quality indicator species on lodgepole pine (Pinus contorta) and white spruce (Picea alba) stands in west-central Alberta [34]. It is also used as a secondary indicator of slope instability in grand fir (Abies grandis)/pachistima (Pachistima myrsinites) habitat types on the Clearwater National Forest, Idaho [27]. SEASONAL DEVELOPMENT : Oak fern unfolds its fronds in early spring [6] and senesces in autumn [16].

FIRE ECOLOGY

SPECIES: Gymnocarpium dryopteris | Oak Fern
FIRE ECOLOGY OR ADAPTATIONS : Oak fern has rhizomes which may allow it to sprout following fire [25]. Because spores are stored in the soil seedbank, fires that do not damage upper soil layers may not permanently eliminate oak fern from an area. POSTFIRE REGENERATION STRATEGY : Rhizomatous herb, rhizome in soil

FIRE EFFECTS

SPECIES: Gymnocarpium dryopteris | Oak Fern
IMMEDIATE FIRE EFFECT ON PLANT : Fire can topkill oak fern, and repeated burning can significantly reduce it's frequency [25,32]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : In white spruce climax forests of Alaska, light surface fires usually do not affect understory species composition, of which oak fern is a part [22]. However, stand-replacement fires that completely eliminate white spruce result in early seral communities where oak fern is not present. In cedar-hemlock forests of northern Idaho, oak fern successively decreased in abundance on sites that were logged, slashpile-burned, broadcast burned once, and burned two or more times over a 30-year period [25]. PLANT RESPONSE TO FIRE : Oak fern appears to decrease in constancy and/or cover following logging and burning [14]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : In the East Slope Region of central Alberta, oak fern is not present until the climax stages of succession following fire [9]. On severely burned sites in northern Idaho (where all trees and groundlayer vegetation was consumed), oak fern appeared in the third postfire year only [32]. This occurence was rare because oak fern is not considered a fire-surviving species. In the subboreal spruce zone of British Columbia, oak fern was present within 10 years following fire on four sites ranging from fairly dry to wet [14]. Fires were broadcast burns following logging, and its effects on specific plants were not studied at the time of the fire. FIRE MANAGEMENT CONSIDERATIONS : NO-ENTRY

REFERENCES

SPECIES: Gymnocarpium dryopteris | Oak Fern
REFERENCES : 1. Alaback, Paul B. 1982. Dynamics of understory biomass in Sitka spruce-western hemlock forests of southeast Alaska. Ecology. 63(6): 1932-1948. [7305] 2. Almack, Jon. 1986. Grizzly bear habitat use, food habits, and movements in the Selkirk Mountains, northern Idaho. In: Contreras, Glen P.; Evans, Keith E., compilers. Proceedings--grizzly bear habitat symposium; 1985 April 30 - May 2; Missoula, MT. Gen. Tech. Rep. INT-207. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 150-157. [10815] 3. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434] 4. Boggs, Keith; Hansen, Paul; Pfister, Robert; Joy, John. 1990. Classification and management of riparian and wetland sites in northwestern Montana. Missoula, MT: University of Montana, School of Forestry, Montana Forest and Conservation Experiment Station, Montana Riparian Association. 217 p. Draft Version 1. [8447] 5. Caza, C, L.; Kimmins, J. P. 1990. Problems with the development and application of competition indices in complex, multispecies communities. In: Hamilton, Evelyn, compiler. Vegetation management: An integrated approach--Proceedings of the 4th annual vegetation management workshop; 1989 November 14-16; Vancouver, BC. FRDA Report 109. Victoria, BC: Ministry of Forests, Research Branch: 30-32. [10950] 6. Cody, William J.; Britton, Donald M. 1989. Ferns and fern allies of Canada. Ottawa, ON: Agriculture Canada, Research Branch. 430 p. [13078] 7. Cooper, Stephen V.; Neiman, Kenneth E.; Roberts, David W. 1991. (Rev.) Forest habitat types of northern Idaho: a second approximation. Gen. Tech. Rep. INT-236. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 143 p. [14792] 8. Cormack, R. G. H. 1949. A study of trout streamside cover in logged-over and undisturbed virgin spruce woods. Canadian Journal of Research. 27: 78-95. [15689] 9. Cormack, R. G. H. 1953. A survey of coniferous forest succession in the eastern Rockies. Forestry Chronicle. 29: 218-232. [16458] 10. Corns, I. G. W.; Annas, R. M. 1986. Field guide to forest ecosystems of west-central Alberta. Edmonton, AB: Canadian Forestry Service, Northern Forestry Centre. 251 p. [8998] 11. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 12. Franklin, Jerry F.; Moir, William H.; Hemstrom, Miles A.; [and others]. 1988. The forest communities of Mount Rainier National Park. Scientific Monograph Series No 19. Washington, DC: U.S. Department of the Interior, National Park Service. 194 p. [12392] 13. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998] 14. Hamilton, Evelyn H.; Yearsley, H. Karen. 1988. Vegetation development after clearcutting and site preparation in the SBS zone. Economic and Regional Development Agreement: FRDA Report 018. Victoria, BC: Canadian Forestry Service, Pacific Forestry Centre; British Columbia Ministry of Forests and Lands. 66 p. [8760] 15. Harcombe, Andrew; Pendergast, Bruce; Petch, Bruce; Janz, Doug. 1983. Elk habitat management: Salmon River Valley. MOE Working Report 1. 83-05-10. Victoria, BC: Ministry of the Environment. 83 p. [9984] 16. Hitchcock, C. Leo; Cronquist, Arthur; Ownbey, Marion. 1969. Vascular plants of the Pacific Northwest. Part 1: Vascular cryptograms, gymnosperms, and monocotyledons. Seattle, WA: University of Washington Press. 914 p. [1169] 17. Jones, R. Keith; Pierpoint, Geoffrey; Wickware, Gregory M.; [and others]. 1983. Field guide to forest ecosystem classification for the Clay Belt, site region 3e. Maple, Ontario: Ministry of Natural Resources, Ontario Forest Research Institute. 160 p. [16163] 18. Kirkpatrick, Ruth E. B.; Soltis, Pamela S.; Soltis, Douglas E. 1990. Mating system and distribution of genetic variation in Gymnocarpium dryopteris ssp. disjunctum. American Journal of Botany. 77(8): 1101-1110. [16231] 19. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384] 20. Kudish, Michael. 1992. Adirondack upland flora: an ecological perspective. Saranac, NY: The Chauncy Press. 320 p. [19376] 21. Lackschewitz, Klaus. 1991. Vascular plants of west-central Montana--identification guidebook. Gen. Tech. Rep. INT-227. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 648 p. [13798] 22. Lutz, H. J. 1953. The effects of forest fires on the vegetation of interior Alaska. Juneau, AK: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 36 p. [7076] 23. Milberg, Per. 1991. Fern spores in a grassland soil. Canadian Journal of Botany. 69: 831-834. [14868] 24. Mladenoff, David J. 1990. The relationship of the soil seed bank and understory vegetation in old-growth northern hardwood-hemlock treefall gaps. Canadian Journal of Botany. 68: 2714-2721. [13477] 25. Mueggler, Walter F. 1965. Ecology of seral shrub communities in the cedar-hemlock zone of northern Idaho. Ecological Monographs. 35: 165-185. [4016] 26. Otchere-Boateng, J.; Herring, L. J. 1990. Site preparation: chemical. In: Lavender, D. P.; Parish, R.; Johnson, C. M.; [and others], eds. Regenerating British Columbia's Forests. Vancouver, BC: University of British Columbia Press: 164-178. [10714] 27. Pole, Michael W.; Satterlund, Donald R. 1978. Plant indicators of slope instability. Journal of Soil and Water Conservation. Sept-Oct: 230-232. [8199] 28. Pryer, Kathleen M.; Haufler, Christopher H. 1993. Isozymic and chromosomal evidence for the allotetraploid origin of Gymnocarpium dryopteris (Dryopteridaceae). Systematic Botany. 18(1): 150-172. [21783] 29. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 30. Reynolds, Keith M. 1990. Preliminary classification of forest vegetation of the Kenai Penninsula, Alaska. Res. Pap. PNW-RP-424. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 67 p. [14581] 31. Shearer, Raymond C.; Stickney, Peter F. 1991. Natural revegetation of burned and unburned clearcuts in western larch forests of northwest Montana. In: Nodvin, Stephen C.; Waldrop, Thomas A., eds. Fire and the environment: ecological and cultural perspectives: Proceedings of an international symposium; 1990 March 20-24; Knoxville, TN. Gen. Tech. Rep. SE-69. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 66-74. [16635] 32. Stickney, Peter F. 1986. First decade plant succession following the Sundance Forest Fire, northern Idaho. Gen. Tech. Rep. INT-197. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 26 p. [2255] 33. Stickney, Peter F. 1989. Seral origin of species originating in northern Rocky Mountain forests. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT; RWU 4403 files. 7 p. [20090] 34. Strong, W. L.; Pluth, D. J.; LaRoi, G. H.; Corns, I. G. W. 1991. Forest understory plants as predictors of lodgepole pine and white spruce site quality in west-central Alberta. Canadian Journal of Forest Research. 21: 1675-1683. [17695] 35. U.S. Department of Agriculture, Soil Conservation Service. 1982. National list of scientific plant names. Vol. 1. List of plant names. SCS-TP-159. Washington, DC. 416 p. [11573] 36. Zoladeski, C. A. 1988. Classification and gradient analysis of forest vegetation of Cape Enrage, Bic Park, Quebec. Le Naturaliste Canadien. 115(1): 9-18. [13610]

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